Bacteria

Bacteria
Primates
Bacteria
Flowers
Birds Mammals listen); common noun bacteria, singular
Plants Dinosaurs biological
Bacteria
bacterium cell. They constitute a large domain
of microorganisms. Typically a few micrometres in length,
Temporal range:
ArthropodsMolluscs
bacteria have a number of shapes, ranging from spheres to rods and Archean or earlier – present
spirals. Bacteria were among the first life forms to appear on Earth, Had'n Archean Proterozoic Pha.
and areMulticellular
present in most of its habitats. Bacteria inhabit soil, water,
life radioactive waste,[3] and the deep portions of Earth's
acidic hot springs
crust. Bacteria also live in symbiotic and parasitic relationships with
plants and animals. Most bacteria have not been characterised, and
only about half of the bacterial phyla have species that can be grown in
the laboratory. The study of bacteria is known as bacteriology, a
Eukaryotes
branch of
There are typically 40 million bacterial cells in a gram of soil and a

million bacterial cells in a millilitre of fresh water. There are
Scanning electron micrograph of
approximately 5×10 bacteria on Earth,[5] forming a biomass which
exceeds that of all plants and animals.[6] Bacteria are vital in many
Escherichia coli rods
photosynthesis
stages of the nutrient cycle by recycling nutrients such as the fixation Scientific classification
of nitrogen atmosphere. The nutrient cycle includes the
Domain: Bacteria
decomposition dead bodies and bacteria are responsible for the
Woese, Kandler & Wheelis,
putrefaction stage in this process.[7] In the biological communities
1990[1]
surrounding hydrothermal vents and cold seeps, extremophile bacteria
provide the nutrients needed to sustain life by converting dissolved
Single-celled Phyla
compounds, such as hydrogen sulphide and methane, to energy. In
life Acidobacteria
March 2013, data reported by researchers in October 2012, was
Actinobacteria
published. It was suggested that bacteria thrive in the Mariana Trench,
water Aquificae
which with a depth of up to 11 kilometres is the deepest known part of
Armatimonadetes
the oceans.[8][9] Other researchers reported related studies that
Bacteroidetes
microbes thrive inside rocks up to 580 metres below the sea floor
Caldiserica
under 2.6 kilometres of ocean off the coast of the northwestern United
Chlamydiae
States.[8][10] According to one of the researchers, "You can find
Chlorobi
microbes everywhere—they're extremely adaptable to conditions, and
Chloroflexi
survive wherever they are."[8]
Chrysiogenetes
The famous notion that bacterial cells in the human body outnumber Cyanobacteria
human cells by a factor of 10:1 has been debunked. There are Deferribacteres
approximately 39 trillion bacterial cells in the human microbiota as Deinococcus-Thermus
personified by a "reference" 70 kg male 170 cm tall, whereas there are Dictyoglomi
30 trillion human cells in the body. This means that although they do Elusimicrobia
have the upper hand in actual numbers, it is only by 30%, and not Fibrobacteres
900%.[11] Firmicutes
6/4/2018, 9:29 AM 1
The largest number exist in the gut flora, and a large number on the Fusobacteria
skin.[12] The vast majority of the bacteria in the body are rendered Gemmatimonadetes
harmless by the protective effects of the immune system, though many Lentisphaerae
are beneficial, particularly in the gut flora. However several species of Nitrospirae
bacteria are pathogenic and cause infectious diseases, including Planctomycetes
cholera, syphilis, anthrax, leprosy, and bubonic plague. The most Proteobacteria
common fatal bacterial diseases are respiratory infections, with Spirochaetes
tuberculosis alone killing about 2 million people per year, mostly in Synergistetes
sub-Saharan Africa.[13] In developed countries, antibiotics are used to Tenericutes
treat bacterial infections and are also used in farming, making Thermodesulfobacteria
antibiotic resistance a growing problem. In industry, bacteria are Thermotogae
important in sewage treatment and the breakdown of oil spills, the Verrucomicrobia
production of cheese and yogurt through fermentation, the recovery of
gold, palladium, copper and other metals in the mining sector,[14] as Synonyms
well as in biotechnology, and the manufacture of antibiotics and other
Eubacteria Woese & Fox, 1977[2]
chemicals.[15]
Once regarded as plants constituting the class Schizomycetes, bacteria

are now classified as prokaryotes. Unlike cells of animals and other eukaryotes, bacterial cells do not contain a
nucleus and rarely harbour membrane-bound organelles. Although the term bacteria traditionally included all
prokaryotes, the scientific classification changed after the discovery in the 1990s that prokaryotes consist of two very
different groups of organisms that evolved from an ancient common ancestor. These evolutionary domains are called
Bacteria and Archaea.[1]
Contents
Etymology
Origin and early evolution
Morphology
Cellular structure
Intracellular structures
Extracellular structures
Endospores
Metabolism
Growth and reproduction
Genetics
Behaviour
Movement
Communication
Classification and identification

Interactions with other organisms
Predators
6/4/2018, 9:29 AM 2
Mutualists
Pathogens
Significance in technology and industry

History of bacteriology
See also
References
Further reading
External links
Etymology
The word bacteria is the plural of the New Latin
bacterium, which is the latinisation of the Greek Life timeline
βακτήριον (bakterion),[16] the diminutive of view • discuss •
ice ages
βακτηρία (bakteria), meaning "staff, cane",[17] 0—
Quaternary P
←Earliest apes
because the first ones to be discovered were rod- h
shaped.[18][19] Karoo
– a
n
Andean e ←Tetrapoda
-500 — r ←Cambrian explosion
o ←Ediacara biota
Origin and early Cryogenian
–
z
o
←Earliest plants
evolution -1000 —
i
c
The ancestors of modern bacteria were unicellular

–
←Earliest sexual
microorganisms that were the first forms of life to
reproduction
P
-1500 — r
appear on Earth, about 4 billion years ago. For o
t
about 3 billion years, most organisms were e
–
r
microscopic, and bacteria and archaea were the o
dominant forms of life.[20][21] Although bacterial -2000 — z
o
fossils exist, such as stromatolites, their lack of i
Huronian – c ←Oxygen crisis
distinctive morphology prevents them from being
used to examine the history of bacterial evolution, -2500 — ←Atmospheric oxygen
or to date the time of origin of a particular bacterial –
species. However, gene sequences can be used to Pongola
reconstruct the bacterial phylogeny, and these -3000 — A

r
studies indicate that bacteria diverged first from the c
– h
archaeal/eukaryotic lineage.[22] The most recent e
a
common ancestor of bacteria and archaea was -3500 — n ←Earliest oxygen
probably a hyperthermophile that lived about 2.5 –
billion–3.2 billion years ago.[23][24] ←LHB meteorites
-4000 —
H
Bacteria were also involved in the second great a
– d ←Earliest life
evolutionary divergence, that of the archaea and e
a ←Earliest water
eukaryotes. Here, eukaryotes resulted from the -4500 — n ←Earliest Earth
6/4/2018, 9:29 AM 3
(−4540)
entering of ancient bacteria into endosymbiotic Axis scale: million years
associations with the ancestors of eukaryotic cells, Also see: Human timeline and Nature timeline
which were themselves possibly related to the
Archaea.[25][26] This involved the engulfment by proto-eukaryotic cells of alphaproteobacterial symbionts to form
either mitochondria or hydrogenosomes, which are still found in all known Eukarya (sometimes in highly reduced
form, e.g. in ancient "amitochondrial" protozoa). Later, some eukaryotes that already contained mitochondria also
engulfed cyanobacteria-like organisms, leading to the formation of chloroplasts in algae and plants.[27][28] This is
known as secondary endosymbiosis.
Morphology
Bacteria display a wide diversity of shapes and sizes,
called morphologies. Bacterial cells are about one-
tenth the size of eukaryotic cells and are typically
0.5–5.0 micrometres in length. However, a few
species are visible to the unaided eye—for example,
Thiomargarita namibiensis is up to half a
millimetre long[29] and Epulopiscium fishelsoni
reaches 0.7 mm.[30] Among the smallest bacteria are
members of the genus Mycoplasma, which measure
only 0.3 micrometres, as small as the largest
viruses.[31] Some bacteria may be even smaller, but
these ultramicrobacteria are not well-studied.[32]
Most bacterial species are either spherical, called

cocci (sing. coccus, from Greek kókkos, grain, seed),
or rod-shaped, called bacilli (sing. bacillus, from
Bacteria display many cell morphologies and arrangements Latin baculus, stick).[33] Some bacteria, called vibrio,
are shaped like slightly curved rods or comma-
shaped; others can be spiral-shaped, called spirilla,
or tightly coiled, called spirochaetes. A small number of other unusual shapes have been described, such as star-
shaped bacteria.[34] This wide variety of shapes is determined by the bacterial cell wall and cytoskeleton, and is
important because it can influence the ability of bacteria to acquire nutrients, attach to surfaces, swim through
liquids and escape predators.[35][36]
Many bacterial species exist simply as single cells, others associate in characteristic patterns: Neisseria form diploids
(pairs), Streptococcus form chains, and Staphylococcus group together in "bunch of grapes" clusters. Bacteria can
also group to form larger multicellular structures, such as the elongated filaments of Actinobacteria, the aggregates
of Myxobacteria, and the complex hyphae of Streptomyces.[37] These multicellular structures are often only seen in
certain conditions. For example, when starved of amino acids, Myxobacteria detect surrounding cells in a process
known as quorum sensing, migrate towards each other, and aggregate to form fruiting bodies up to 500 micrometres
long and containing approximately 100,000 bacterial cells.[38] In these fruiting bodies, the bacteria perform separate
tasks; for example, about one in ten cells migrate to the top of a fruiting body and differentiate into a specialised
dormant state called a myxospore, which is more resistant to drying and other adverse environmental conditions.[39]
Bacteria often attach to surfaces and form dense aggregations called biofilms, and larger formations known as
6/4/2018, 9:29 AM 4
microbial mats. These biofilms and mats can range from a few
micrometres in thickness to up to half a metre in depth, and may contain
multiple species of bacteria, protists and archaea. Bacteria living in
biofilms display a complex arrangement of cells and extracellular
components, forming secondary structures, such as microcolonies,
through which there are networks of channels to enable better diffusion of
nutrients.[40][41] In natural environments, such as soil or the surfaces of
plants, the majority of bacteria are bound to surfaces in biofilms.[42]
Biofilms are also important in medicine, as these structures are often
present during chronic bacterial infections or in infections of implanted The range of sizes shown by
medical devices, and bacteria protected within biofilms are much harder to
kill than individual isolated bacteria.[43] prokaryotes, relative to those of
other organisms and biomolecules.
Cellular structure
Intracellular structures
The bacterial cell is surrounded by a cell membrane which is made
primarily of phospholipids. This membrane encloses the contents of the
cell and acts as a barrier to hold nutrients, proteins and other essential
components of the cytoplasm within the cell.[44] Unlike eukaryotic cells,
bacteria usually lack large membrane-bound structures in their cytoplasm
such as a nucleus, mitochondria, chloroplasts and the other organelles
present in eukaryotic cells.[45] However, some bacteria have protein-bound
organelles in the cytoplasm which compartmentalize aspects of bacterial
metabolism,[46][47] such as the carboxysome.[48] Additionally, bacteria have
a multi-component cytoskeleton to control the localisation of proteins and
Structure and contents of a typical
nucleic acids within the cell, and to manage the process of cell division.
[49][50][51]
gram-positive bacterial cell (seen by
the fact that only one cell membrane
is present).
Many important biochemical reactions, such as energy generation, occur
due to concentration gradients across membranes, creating a potential
difference analogous to a battery. The general lack of internal membranes in bacteria means these reactions, such as
electron transport, occur across the cell membrane between the cytoplasm and the outside of the cell or periplasm.[52]
However, in many photosynthetic bacteria the plasma membrane is highly folded and fills most of the cell with layers
of light-gathering membrane.[53] These light-gathering complexes may even form lipid-enclosed structures called
chlorosomes in green sulfur bacteria.[54]
Most bacteria do not have a membrane-bound nucleus, and their genetic material is typically a single circular
bacterial chromosome of DNA located in the cytoplasm in an irregularly shaped body called the nucleoid.[55] The
nucleoid contains the chromosome with its associated proteins and RNA. Like all living organisms, bacteria contain
ribosomes for the production of proteins, but the structure of the bacterial ribosome is different from that of
eukaryotes and Archaea.[56]
Some bacteria produce intracellular nutrient storage granules, such as glycogen,[57] polyphosphate,[58] sulfur[59] or
6/4/2018, 9:29 AM 5
polyhydroxyalkanoates.[60] Certain bacterial species, such as the
photosynthetic Cyanobacteria, produce internal gas vacuoles which they
use to regulate their buoyancy, allowing them to move up or down into
water layers with different light intensities and nutrient levels.[61]
Extracellular structures
Around the outside of the cell membrane is the cell wall. Bacterial cell An electron micrograph of
walls are made of peptidoglycan (called "murein" in older sources), which Halothiobacillus neapolitanus cells
is made from polysaccharide chains cross-linked by peptides containing with carboxysomes inside, with
D-amino acids.[62] Bacterial cell walls are different from the cell walls of arrows highlighting visible
plants and fungi, which are made of cellulose and chitin, respectively.[63] carboxysomes. Scale bars indicate
100 nm.
The cell wall of bacteria is also distinct from that of Archaea, which do not
contain peptidoglycan. The cell wall is essential to the survival of many
bacteria, and the antibiotic penicillin is able to kill bacteria by inhibiting a step in the synthesis of peptidoglycan.[63]
There are broadly speaking two different types of cell wall in bacteria, called Gram-positive and Gram-negative. The
names originate from the reaction of cells to the Gram stain, a long-standing test for the classification of bacterial
species.[64]
Gram-positive bacteria possess a thick cell wall containing many layers of peptidoglycan and teichoic acids. In
contrast, Gram-negative bacteria have a relatively thin cell wall consisting of a few layers of peptidoglycan
surrounded by a second lipid membrane containing lipopolysaccharides and lipoproteins. Most bacteria have the
Gram-negative cell wall, and only the Firmicutes and Actinobacteria (previously known as the low G+C and high G+C
gram-positive bacteria, respectively) have the alternative Gram-positive arrangement.[65] These differences in
structure can produce differences in antibiotic susceptibility; for instance, vancomycin can kill only Gram-positive
bacteria and is ineffective against Gram-negative pathogens, such as Haemophilus influenzae or Pseudomonas
aeruginosa.[66] Some bacteria have cell wall structures that are neither classically Gram-positive or Gram-negative.
This includes clinically important bacteria such as Mycobacteria which have a thick peptidoglycan cell wall like a
Gram-positive bacterium, but also a second outer layer of lipids.[67]
In many bacteria, an S-layer of rigidly arrayed protein molecules covers the outside of the cell.[68] This layer provides
chemical and physical protection for the cell surface and can act as a macromolecular diffusion barrier. S-layers have
diverse but mostly poorly understood functions, but are known to act as virulence factors in Campylobacter and
contain surface enzymes in Bacillus stearothermophilus.[69]
Flagella are rigid protein structures, about 20 nanometres in diameter and up to 20 micrometres in length, that are
used for motility. Flagella are driven by the energy released by the transfer of ions down an electrochemical gradient
across the cell membrane.[70]
Fimbriae (sometimes called "attachment pili") are fine filaments of protein, usually 2–10 nanometres in diameter
and up to several micrometres in length. They are distributed over the surface of the cell, and resemble fine hairs
when seen under the electron microscope. Fimbriae are believed to be involved in attachment to solid surfaces or to
other cells, and are essential for the virulence of some bacterial pathogens.[71] Pili (sing. pilus) are cellular
appendages, slightly larger than fimbriae, that can transfer genetic material between bacterial cells in a process called
conjugation where they are called conjugation pili or sex pili (see bacterial genetics, below).[72] They can also generate
6/4/2018, 9:29 AM 6
movement where they are called type IV pili.[73]
Glycocalyx is produced by many bacteria to surround their cells, and varies

in structural complexity: ranging from a disorganised slime layer of
extracellular polymeric substances to a highly structured capsule. These
structures can protect cells from engulfment by eukaryotic cells such as
macrophages (part of the human immune system).[74] They can also act as
antigens and be involved in cell recognition, as well as aiding attachment
Helicobacter pylori electron to surfaces and the formation of biofilms.[75]
micrograph, showing multiple flagella
on the cell surface The assembly of these extracellular structures is dependent on bacterial
secretion systems. These transfer proteins from the cytoplasm into the
periplasm or into the environment around the cell. Many types of
secretion systems are known and these structures are often essential for the virulence of pathogens, so are
intensively studied.[76]
Endospores
Certain genera of Gram-positive bacteria, such as Bacillus, Clostridium,
Sporohalobacter, Anaerobacter, and Heliobacterium, can form highly
resistant, dormant structures called endospores.[77] Endospores develop
within the cytoplasm of the cell; generally a single endospore develops in
each cell.[78] Each endospore contains a core of DNA and ribosomes
surrounded by a cortex layer and protected by a multilayer rigid coat
composed of peptidoglycan and a variety of proteins.[78]
Endospores show no detectable metabolism and can survive extreme Bacillus anthracis (stained purple)
physical and chemical stresses, such as high levels of UV light, gamma growing in cerebrospinal fluid
radiation, detergents, disinfectants, heat, freezing, pressure, and

desiccation.[79] In this dormant state, these organisms may remain viable for millions of years,[80][81] and endospores
even allow bacteria to survive exposure to the vacuum and radiation in space.[82] Endospore-forming bacteria can also
cause disease: for example, anthrax can be contracted by the inhalation of Bacillus anthracis endospores, and
contamination of deep puncture wounds with Clostridium tetani endospores causes tetanus.[83]
Metabolism
Bacteria exhibit an extremely wide variety of metabolic types.[84] The distribution of metabolic traits within a group of
bacteria has traditionally been used to define their taxonomy, but these traits often do not correspond with modern
genetic classifications.[85] Bacterial metabolism is classified into nutritional groups on the basis of three major
criteria: the source of energy, the electron donors used, and the source of carbon used for growth.[86]
Bacteria either derive energy from light using photosynthesis (called phototrophy), or by breaking down chemical
compounds using oxidation (called chemotrophy).[87] Chemotrophs use chemical compounds as a source of energy by
transferring electrons from a given electron donor to a terminal electron acceptor in a redox reaction. This reaction
releases energy that can be used to drive metabolism. Chemotrophs are further divided by the types of compounds
they use to transfer electrons. Bacteria that use inorganic compounds such as hydrogren, carbon monoxide, or
6/4/2018, 9:29 AM 7
ammonia as sources of electrons are called lithotrophs, while those that use organic compounds are called
organotrophs.[87] The compounds used to receive electrons are also used to classify bacteria: aerobic organisms use
oxygen as the terminal electron acceptor, while anaerobic organisms use other compounds such as nitrate, sulfate, or
carbon dioxide.[87]
Many bacteria get their carbon from other organic carbon, called heterotrophy. Others such as cyanobacteria and
some purple bacteria are autotrophic, meaning that they obtain cellular carbon by fixing carbon dioxide.[88] In
unusual circumstances, the gas methane can be used by methanotrophic bacteria as both a source of electrons and a
substrate for carbon anabolism.[89]
Nutritional types in bacterial metabolism
Nutritional Source of
Source of carbon Examples
type energy
Organic compounds
Cyanobacteria, Green sulfur bacteria,
Phototrophs Sunlight (photoheterotrophs) or carbon fixation
Chloroflexi, or Purple bacteria
(photoautotrophs)
Inorganic Organic compounds (lithoheterotrophs) Thermodesulfobacteria,

Lithotrophs
compounds or carbon fixation (lithoautotrophs) Hydrogenophilaceae, or Nitrospirae
Organic compounds
Organic Bacillus, Clostridium or
Organotrophs (chemoheterotrophs) or carbon fixation
compounds Enterobacteriaceae
(chemoautotrophs)
In many ways, bacterial metabolism provides traits that are useful for ecological stability and for human society. One
example is that some bacteria have the ability to fix nitrogen gas using the enzyme nitrogenase. This environmentally
important trait can be found in bacteria of most metabolic types listed above.[90] This leads to the ecologically
important processes of denitrification, sulfate reduction, and acetogenesis, respectively.[91][92] Bacterial metabolic
processes are also important in biological responses to pollution; for example, sulfate-reducing bacteria are largely
responsible for the production of the highly toxic forms of mercury (methyl- and dimethylmercury) in the
environment.[93] Non-respiratory anaerobes use fermentation to generate energy and reducing power, secreting
metabolic by-products (such as ethanol in brewing) as waste. Facultative anaerobes can switch between fermentation
and different terminal electron acceptors depending on the environmental conditions in which they find themselves.
Growth and reproduction

Unlike in multicellular organisms, increases in cell size (cell growth) and reproduction by cell division are tightly
linked in unicellular organisms. Bacteria grow to a fixed size and then reproduce through binary fission, a form of
asexual reproduction.[94] Under optimal conditions, bacteria can grow and divide extremely rapidly, and bacterial
populations can double as quickly as every 9.8 minutes.[95] In cell division, two identical clone daughter cells are
produced. Some bacteria, while still reproducing asexually, form more complex reproductive structures that help
disperse the newly formed daughter cells. Examples include fruiting body formation by Myxobacteria and aerial
hyphae formation by Streptomyces, or budding. Budding involves a cell forming a protrusion that breaks away and
produces a daughter cell.
In the laboratory, bacteria are usually grown using solid or liquid media. Solid growth media, such as agar plates, are
used to isolate pure cultures of a bacterial strain. However, liquid growth media are used when measurement of
growth or large volumes of cells are required. Growth in stirred liquid media occurs as an even cell suspension,
6/4/2018, 9:29 AM 8
making the cultures easy to divide and
transfer, although isolating single bacteria
from liquid media is difficult. The use of
selective media (media with specific nutrients
added or deficient, or with antibiotics added)
can help identify specific organisms.[97]
A colony of
Escherichia Most laboratory techniques for growing
coli [96] bacteria use high levels of nutrients to produce
large amounts of cells cheaply and quickly.
However, in natural environments, nutrients
are limited, meaning that bacteria cannot continue to reproduce
indefinitely. This nutrient limitation has led the evolution of different
growth strategies (see r/K selection theory). Some organisms can Many bacteria reproduce through binary
grow extremely rapidly when nutrients become available, such as the fission, which is compared to mitosis and
meiosis in this image.
formation of algal (and cyanobacterial) blooms that often occur in
lakes during the summer.[98] Other organisms have adaptations to
harsh environments, such as the production of multiple antibiotics by Streptomyces that inhibit the growth of
competing microorganisms.[99] In nature, many organisms live in communities (e.g., biofilms) that may allow for
increased supply of nutrients and protection from environmental stresses.[42] These relationships can be essential for
growth of a particular organism or group of organisms (syntrophy).[100]
Bacterial growth follows four phases. When a population of bacteria first enter a high-nutrient environment that
allows growth, the cells need to adapt to their new environment. The first phase of growth is the lag phase, a period of
slow growth when the cells are adapting to the high-nutrient environment and preparing for fast growth. The lag
phase has high biosynthesis rates, as proteins necessary for rapid growth are produced.[101] The second phase of
growth is the logarithmic phase, also known as the exponential phase. The log phase is marked by rapid exponential
growth. The rate at which cells grow during this phase is known as the growth rate (k), and the time it takes the cells
to double is known as the generation time (g). During log phase, nutrients are metabolised at maximum speed until
one of the nutrients is depleted and starts limiting growth. The third phase of growth is the stationary phase and is
caused by depleted nutrients. The cells reduce their metabolic activity and consume non-essential cellular proteins.
The stationary phase is a transition from rapid growth to a stress response state and there is increased expression of
genes involved in DNA repair, antioxidant metabolism and nutrient transport.[102] The final phase is the death phase
where the bacteria run out of nutrients and die.[103]
Genetics
Most bacteria have a single circular chromosome that can range in size from only 160,000 base pairs in the
endosymbiotic bacteria Carsonella ruddii,[104] to 12,200,000 base pairs (12.2 Mbp) in the soil-dwelling bacteria
Sorangium cellulosum.[105] There are many exceptions to this, for example some Streptomyces and Borrelia species
contain a single linear chromosome,[106][107] while some Vibrio species contain more than one chromosome.[108]
Bacteria can also contain plasmids, small extra-chromosomal DNAs that may contain genes for various useful
functions such as antibiotic resistance, metabolic capabilities, or various virulence factors.[109]
Bacteria genomes usually encode a few hundred to a few thousand genes. The genes in bacterial genomes are usually
a single continuous stretch of DNA and although several different types of introns do exist in bacteria, these are
6/4/2018, 9:29 AM 9
much rarer than in eukaryotes.[110]
Bacteria, as asexual organisms, inherit an identical copy of the parent's genomes and are clonal. However, all bacteria
can evolve by selection on changes to their genetic material DNA caused by genetic recombination or mutations.
Mutations come from errors made during the replication of DNA or from exposure to mutagens. Mutation rates vary
widely among different species of bacteria and even among different clones of a single species of bacteria.[111] Genetic
changes in bacterial genomes come from either random mutation during replication or "stress-directed mutation",
where genes involved in a particular growth-limiting process have an increased mutation rate.[112]
Some bacteria also transfer genetic material between cells. This can occur in three main ways. First, bacteria can take
up exogenous DNA from their environment, in a process called transformation.[113] Many bacteria can naturally take
up DNA from the environment, while others must be chemically altered in order to induce them to take up DNA.[114]
The development of competence in nature is usually associated with stressful environmental conditions, and seems
to be an adaptation for facilitating repair of DNA damage in recipient cells.[115] The second way bacteria transfer
genetic material is by transduction, when the integration of a bacteriophage introduces foreign DNA into the
chromosome. Many types of bacteriophage exist, some simply infect and lyse their host bacteria, while others insert
into the bacterial chromosome.[116] Bacteria resist phage infection through restriction modification systems that
degrade foreign DNA,[117] and a system that uses CRISPR sequences to retain fragments of the genomes of phage that
the bacteria have come into contact with in the past, which allows them to block virus replication through a form of
RNA interference.[118][119] The third method of gene transfer is conjugation, whereby DNA is transferred through
direct cell contact. In ordinary circumstances, transduction, conjugation, and transformation involve transfer of DNA
between individual bacteria of the same species, but occasionally transfer may occur between individuals of different
bacterial species and this may have significant consequences, such as the transfer of antibiotic resistance.[120][121] In
such cases, gene acquisition from other bacteria or the environment is called horizontal gene transfer and may be
common under natural conditions.[122]
Behaviour
Movement
Many bacteria are motile and can move using a variety of mechanisms. The best studied of these are flagella, long
filaments that are turned by a motor at the base to generate propeller-like movement.[123] The bacterial flagellum is
made of about 20 proteins, with approximately another 30 proteins required for its regulation and assembly.[123] The
flagellum is a rotating structure driven by a reversible motor at the base that uses the electrochemical gradient across
the membrane for power.[124]
Bacteria can use flagella in different ways to generate different kinds of movement. Many bacteria (such as E. coli)
have two distinct modes of movement: forward movement (swimming) and tumbling. The tumbling allows them to
reorient and makes their movement a three-dimensional random walk.[125] Bacterial species differ in the number and
arrangement of flagella on their surface; some have a single flagellum (monotrichous), a flagellum at each end
(amphitrichous), clusters of flagella at the poles of the cell (lophotrichous), while others have flagella distributed
over the entire surface of the cell (peritrichous). The flagella of a unique group of bacteria, the spirochaetes, are
found between two membranes in the periplasmic space. They have a distinctive helical body that twists about as it
moves.[123]
Two other types of bacterial motion, called twitching motility and gliding motility, rely on a structure called the type
6/4/2018, 9:29 AM 10
IV pilus.[126] In these types of motility, the rod-like pilus extends out from
the cell, binds some substrate, and then retracts, pulling the cell
forward.[127]
Motile bacteria are attracted or repelled by certain stimuli in behaviours

called taxes: these include chemotaxis, phototaxis, energy taxis, and
magnetotaxis.[128][129][130] In one peculiar group, the myxobacteria,
individual bacteria move together to form waves of cells that then
differentiate to form fruiting bodies containing spores.[39] The
myxobacteria move only when on solid surfaces, unlike E. coli, which is
motile in liquid or solid media. Transmission electron micrograph of
Desulfovibrio vulgaris showing a
Several Listeria and Shigella species move inside host cells by usurping
single flagellum at one end of the
the cytoskeleton, which is normally used to move organelles inside the cell. Scale bar is 0.5 micrometers
cell. By promoting actin polymerisation at one pole of their cells, they can long.
form a kind of tail that pushes them through the host cell's cytoplasm.[131]
Communication
A few bacteria have chemical systems that generate light. This
bioluminescence often occurs in bacteria that live in association with fish,
and the light probably serves to attract fish or other large animals.[132]
Bacteria often function as multicellular aggregates known as biofilms,

exchanging a variety of molecular signals for inter-cell communication,
and engaging in coordinated multicellular behaviour.[133][134]
The communal benefits of multicellular cooperation include a cellular

division of labour, accessing resources that cannot effectively be used by
single cells, collectively defending against antagonists, and optimising
population survival by differentiating into distinct cell types.[133] For The different arrangements of
example, bacteria in biofilms can have more than 500 times increased bacterial flagella: A-Monotrichous;
resistance to antibacterial agents than individual "planktonic" bacteria of B-Lophotrichous; C-Amphitrichous;
D-Peritrichous
the same species.[134]
One type of inter-cellular communication by a molecular signal is called

quorum sensing, which serves the purpose of determining whether there is a local population density that is
sufficiently high that it is productive to invest in processes that are only successful if large numbers of similar
organisms behave similarly, as in excreting digestive enzymes or emitting light.
Quorum sensing allows bacteria to coordinate gene expression, and enables them to produce, release and detect
autoinducers or pheromones which accumulate with the growth in cell population.[135]
Classification and identification
Classification seeks to describe the diversity of bacterial species by naming and grouping organisms based on
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similarities. Bacteria can be classified on the basis of cell structure,
cellular metabolism or on differences in cell components, such as DNA,
fatty acids, pigments, antigens and quinones.[97] While these schemes
allowed the identification and classification of bacterial strains, it was
unclear whether these differences represented variation between distinct
species or between strains of the same species. This uncertainty was due
to the lack of distinctive structures in most bacteria, as well as lateral gene
transfer between unrelated species.[137] Due to lateral gene transfer, some
closely related bacteria can have very different morphologies and
Streptococcus mutans visualised
metabolisms. To overcome this uncertainty, modern bacterial with a Gram stain
classification emphasises molecular systematics, using genetic techniques
such as guanine cytosine ratio
determination, genome-genome
hybridisation, as well as sequencing genes
that have not undergone extensive lateral
gene transfer, such as the rRNA gene.[138]
Classification of bacteria is determined by
publication in the International Journal of
Systematic Bacteriology,[139] and Bergey's
Manual of Systematic Bacteriology.[140] The
International Committee on Systematic
Bacteriology (ICSB) maintains international
rules for the naming of bacteria and
taxonomic categories and for the ranking of
them in the International Code of
Phylogenetic tree showing the diversity of bacteria, compared to
Nomenclature of Bacteria.
other organisms.[136] Eukaryotes are coloured red, archaea green
The term "bacteria" was traditionally applied and bacteria blue.
to all microscopic, single-cell prokaryotes.
However, molecular systematics showed
prokaryotic life to consist of two separate domains, originally called Eubacteria and Archaebacteria, but now called
Bacteria and Archaea that evolved independently from an ancient common ancestor.[1] The archaea and eukaryotes
are more closely related to each other than either is to the bacteria. These two domains, along with Eukarya, are the
basis of the three-domain system, which is currently the most widely used classification system in microbiology.[141]
However, due to the relatively recent introduction of molecular systematics and a rapid increase in the number of
genome sequences that are available, bacterial classification remains a changing and expanding field.[4][142] For
example, a few biologists argue that the Archaea and Eukaryotes evolved from gram-positive bacteria.[143]
The identification of bacteria in the laboratory is particularly relevant in medicine, where the correct treatment is
determined by the bacterial species causing an infection. Consequently, the need to identify human pathogens was a
major impetus for the development of techniques to identify bacteria.
The Gram stain, developed in 1884 by Hans Christian Gram, characterises bacteria based on the structural
characteristics of their cell walls.[64] The thick layers of peptidoglycan in the "gram-positive" cell wall stain purple,
while the thin "gram-negative" cell wall appears pink. By combining morphology and Gram-staining, most bacteria
can be classified as belonging to one of four groups (gram-positive cocci, gram-positive bacilli, gram-negative cocci
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and gram-negative bacilli). Some organisms are best identified by stains other than the Gram stain, particularly
mycobacteria or Nocardia, which show acid-fastness on Ziehl–Neelsen or similar stains.[144] Other organisms may
need to be identified by their growth in special media, or by other techniques, such as serology.
Culture techniques are designed to promote the growth and identify particular bacteria, while restricting the growth
of the other bacteria in the sample. Often these techniques are designed for specific specimens; for example, a
sputum sample will be treated to identify organisms that cause pneumonia, while stool specimens are cultured on
selective media to identify organisms that cause diarrhoea, while preventing growth of non-pathogenic bacteria.
Specimens that are normally sterile, such as blood, urine or spinal fluid, are cultured under conditions designed to
grow all possible organisms.[97][145] Once a pathogenic organism has been isolated, it can be further characterised by
its morphology, growth patterns (such as aerobic or anaerobic growth), patterns of hemolysis, and staining.
As with bacterial classification, identification of bacteria is increasingly using molecular methods. Diagnostics using
DNA-based tools, such as polymerase chain reaction, are increasingly popular due to their specificity and speed,
compared to culture-based methods.[146] These methods also allow the detection and identification of "viable but
nonculturable" cells that are metabolically active but non-dividing.[147] However, even using these improved
methods, the total number of bacterial species is not known and cannot even be estimated with any certainty.
Following present classification, there are a little less than 9,300 known species of prokaryotes, which includes
bacteria and archaea;[148] but attempts to estimate the true number of bacterial diversity have ranged from 107 to 109
total species—and even these diverse estimates may be off by many orders of magnitude.[149][150]
Interactions with other organisms

Despite their apparent simplicity, bacteria can form complex
associations with other organisms. These symbiotic
associations can be divided into parasitism, mutualism and
commensalism. Due to their small size, commensal bacteria
are ubiquitous and grow on animals and plants exactly as
they will grow on any other surface. However, their growth
can be increased by warmth and sweat, and large populations
of these organisms in humans are the cause of body odour.
Predators
Some species of bacteria kill and then consume other
microorganisms, these species are called predatory
bacteria.[153] These include organisms such as Myxococcus
xanthus, which forms swarms of cells that kill and digest any
Overview of bacterial infections and main species
bacteria they encounter.[154] Other bacterial predators either
involved.[151][152]
attach to their prey in order to digest them and absorb
nutrients, such as Vampirovibrio chlorellavorus,[155] or
invade another cell and multiply inside the cytosol, such as Daptobacter.[156] These predatory bacteria are thought to
have evolved from saprophages that consumed dead microorganisms, through adaptations that allowed them to
entrap and kill other organisms.[157]
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Mutualists
Certain bacteria form close spatial associations that are essential for their survival. One such mutualistic association,
called interspecies hydrogen transfer, occurs between clusters of anaerobic bacteria that consume organic acids, such
as butyric acid or propionic acid, and produce hydrogen, and methanogenic Archaea that consume hydrogen.[158] The
bacteria in this association are unable to consume the organic acids as this reaction produces hydrogen that
accumulates in their surroundings. Only the intimate association with the hydrogen-consuming Archaea keeps the
hydrogen concentration low enough to allow the bacteria to grow.
In soil, microorganisms that reside in the rhizosphere (a zone that includes the root surface and the soil that adheres
to the root after gentle shaking) carry out nitrogen fixation, converting nitrogen gas to nitrogenous compounds.[159]
This serves to provide an easily absorbable form of nitrogen for many plants, which cannot fix nitrogen themselves.
Many other bacteria are found as symbionts in humans and other organisms. For example, the presence of over 1,000
bacterial species in the normal human gut flora of the intestines can contribute to gut immunity, synthesise vitamins,
such as folic acid, vitamin K and biotin, convert sugars to lactic acid (see Lactobacillus), as well as fermenting
complex undigestible carbohydrates.[160][161][162] The presence of this gut flora also inhibits the growth of potentially
pathogenic bacteria (usually through competitive exclusion) and these beneficial bacteria are consequently sold as
probiotic dietary supplements.[163]
Pathogens
If bacteria form a parasitic association with other organisms, they are
classed as pathogens. Pathogenic bacteria are a major cause of human
death and disease and cause infections such as tetanus, typhoid fever,
diphtheria, syphilis, cholera, foodborne illness, leprosy and tuberculosis. A
pathogenic cause for a known medical disease may only be discovered
many years after, as was the case with Helicobacter pylori and peptic ulcer
disease. Bacterial diseases are also important in agriculture, with bacteria
causing leaf spot, fire blight and wilts in plants, as well as Johne's disease,
mastitis, salmonella and anthrax in farm animals.
Colour-enhanced scanning electron
Each species of pathogen has a characteristic spectrum of interactions
micrograph showing Salmonella
with its human hosts. Some organisms, such as Staphylococcus or typhimurium (red) invading cultured
Streptococcus, can cause skin infections, pneumonia, meningitis and even human cells
overwhelming sepsis, a systemic inflammatory response producing shock,
massive vasodilation and death.[164] Yet these organisms are also part of
the normal human flora and usually exist on the skin or in the nose without causing any disease at all. Other
organisms invariably cause disease in humans, such as the Rickettsia, which are obligate intracellular parasites able
to grow and reproduce only within the cells of other organisms. One species of Rickettsia causes typhus, while
another causes Rocky Mountain spotted fever. Chlamydia, another phylum of obligate intracellular parasites,
contains species that can cause pneumonia, or urinary tract infection and may be involved in coronary heart
disease.[165] Finally, some species, such as Pseudomonas aeruginosa, Burkholderia cenocepacia, and Mycobacterium
avium, are opportunistic pathogens and cause disease mainly in people suffering from immunosuppression or cystic
fibrosis.[166][167]
Bacterial infections may be treated with antibiotics, which are classified as bacteriocidal if they kill bacteria, or
6/4/2018, 9:29 AM 14
bacteriostatic if they just prevent bacterial growth. There are many types of antibiotics and each class inhibits a
process that is different in the pathogen from that found in the host. An example of how antibiotics produce selective
toxicity are chloramphenicol and puromycin, which inhibit the bacterial ribosome, but not the structurally different
eukaryotic ribosome.[168] Antibiotics are used both in treating human disease and in intensive farming to promote
animal growth, where they may be contributing to the rapid development of antibiotic resistance in bacterial
populations.[169] Infections can be prevented by antiseptic measures such as sterilising the skin prior to piercing it
with the needle of a syringe, and by proper care of indwelling catheters. Surgical and dental instruments are also
sterilised to prevent contamination by bacteria. Disinfectants such as bleach are used to kill bacteria or other
pathogens on surfaces to prevent contamination and further reduce the risk of infection.
Significance in technology and industry

Bacteria, often lactic acid bacteria, such as Lactobacillus and Lactococcus, in combination with yeasts and moulds,
have been used for thousands of years in the preparation of fermented foods, such as cheese, pickles, soy sauce,
sauerkraut, vinegar, wine and yogurt.[170][171]
The ability of bacteria to degrade a variety of organic compounds is remarkable and has been used in waste
processing and bioremediation. Bacteria capable of digesting the hydrocarbons in petroleum are often used to clean
up oil spills.[172] Fertiliser was added to some of the beaches in Prince William Sound in an attempt to promote the
growth of these naturally occurring bacteria after the 1989 Exxon Valdez oil spill. These efforts were effective on
beaches that were not too thickly covered in oil. Bacteria are also used for the bioremediation of industrial toxic
wastes.[173] In the chemical industry, bacteria are most important in the production of enantiomerically pure
chemicals for use as pharmaceuticals or agrichemicals.[174]
Bacteria can also be used in the place of pesticides in the biological pest control. This commonly involves Bacillus
thuringiensis (also called BT), a gram-positive, soil dwelling bacterium. Subspecies of this bacteria are used as a
Lepidopteran-specific insecticides under trade names such as Dipel and Thuricide.[175] Because of their specificity,
these pesticides are regarded as environmentally friendly, with little or no effect on humans, wildlife, pollinators and
most other beneficial insects.[176][177]
Because of their ability to quickly grow and the relative ease with which they can be manipulated, bacteria are the
workhorses for the fields of molecular biology, genetics and biochemistry. By making mutations in bacterial DNA and
examining the resulting phenotypes, scientists can determine the function of genes, enzymes and metabolic pathways
in bacteria, then apply this knowledge to more complex organisms.[178] This aim of understanding the biochemistry
of a cell reaches its most complex expression in the synthesis of huge amounts of enzyme kinetic and gene
expression data into mathematical models of entire organisms. This is achievable in some well-studied bacteria, with
models of Escherichia coli metabolism now being produced and tested.[179][180] This understanding of bacterial
metabolism and genetics allows the use of biotechnology to bioengineer bacteria for the production of therapeutic
proteins, such as insulin, growth factors, or antibodies.[181][182]
Because of their importance for research in general, samples of bacterial strains are isolated and preserved in
Biological Resource Centers. This ensures the availability of the strain to scientists worldwide.
History of bacteriology
Bacteria were first observed by the Dutch microscopist Antonie van Leeuwenhoek in 1676, using a single-lens
6/4/2018, 9:29 AM 15
microscope of his own design.[183] He then published his observations in a
series of letters to the Royal Society of London.[184][185][186] Bacteria were
Leeuwenhoek's most remarkable microscopic discovery. They were just at
the limit of what his simple lenses could make out and, in one of the most
striking hiatuses in the history of science, no one else would see them
again for over a century.[187] His observations had also included
protozoans which he called animalcules, and his findings were looked at
again in the light of the more recent findings of cell theory.
Christian Gottfried Ehrenberg introduced the word "bacterium" in

1828.[188] In fact, his Bacterium was a genus that contained non-spore-
forming rod-shaped bacteria,[189] as opposed to Bacillus, a genus of spore-
forming rod-shaped bacteria defined by Ehrenberg in 1835.[190]
Antonie van Leeuwenhoek, the first

Louis Pasteur demonstrated in 1859 that the growth of microorganisms
microbiologist and the first person to
causes the fermentation process, and that this growth is not due to
observe bacteria using a
spontaneous generation. (Yeasts and moulds, commonly associated with microscope.
fermentation, are not bacteria, but rather fungi.) Along with his
contemporary Robert Koch, Pasteur was an early advocate of the germ
theory of disease.[191]
Robert Koch, a pioneer in medical microbiology, worked on cholera, anthrax and tuberculosis. In his research into
tuberculosis Koch finally proved the germ theory, for which he received a Nobel Prize in 1905.[192] In Koch's
postulates, he set out criteria to test if an organism is the cause of a disease, and these postulates are still used
today.[193]
Ferdinand Cohn is said to be a founder of bacteriology, studying bacteria from 1870. Cohn was the first to classify
bacteria based on their morphology.[194][195]
Though it was known in the nineteenth century that bacteria are the cause of many diseases, no effective
antibacterial treatments were available.[196] In 1910, Paul Ehrlich developed the first antibiotic, by changing dyes that
selectively stained Treponema pallidum—the spirochaete that causes syphilis—into compounds that selectively killed
the pathogen.[197] Ehrlich had been awarded a 1908 Nobel Prize for his work on immunology, and pioneered the use
of stains to detect and identify bacteria, with his work being the basis of the Gram stain and the Ziehl–Neelsen
stain.[198]
A major step forward in the study of bacteria came in 1977 when Carl Woese recognised that archaea have a separate
line of evolutionary descent from bacteria.[2] This new phylogenetic taxonomy depended on the sequencing of 16S
ribosomal RNA, and divided prokaryotes into two evolutionary domains, as part of the three-domain system.[1]
See also
Bacteriotherapy
Genetically modified bacteria
List of bacterial orders
Panspermia
Polysaccharide encapsulated bacteria
6/4/2018, 9:29 AM 16
Psychrotrophic bacteria
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Further reading
Alcamo IE (2001). Fundamentals of microbiology. Boston: Jones and Bartlett. ISBN 0-7637-1067-9.
Atlas RM (1995). Principles of microbiology. St. Louis: Mosby. ISBN 0-8016-7790-4.
Martinko JM, Madigan MT (2005). Brock Biology of Microorganisms (11th ed.). Englewood Cliffs, N.J: Prentice
Hall. ISBN 0-13-144329-1.
Holt JC, Bergey DH (1994). Bergey's manual of determinative bacteriology (9th ed.). Baltimore: Williams &
Wilkins. ISBN 0-683-00603-7.
Hugenholtz P, Goebel BM, Pace NR (September 1998). "Impact of culture-independent studies on the emerging
phylogenetic view of bacterial diversity" (http://jb.asm.org/cgi/content/full/180/18/4765?view=full&pmid=9733676).
Journal of Bacteriology. 180 (18): 4765–74. PMC 107498 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC107498)
. PMID 9733676 (https://www.ncbi.nlm.nih.gov/pubmed/9733676).
Funke BR, Tortora GJ, Case CL (2004). Microbiology: an introduction (8th ed.). San Francisco: Benjamin
Cummings. ISBN 0-8053-7614-3.
Ogunseitan OA (2005). Microbial Diversity: Form and Function in Prokaryotes. Wiley-Blackwell.
ISBN 978-1-4051-4448-3.
Shively JM (2006). Complex Intracellular Structures in Prokaryotes (Microbiology Monographs). Berlin: Springer.
ISBN 3-540-32524-7.
6/4/2018, 9:29 AM 32
External links
MicrobeWiki (http://microbewiki.kenyon.edu/index.php/MicrobeWiki), an extensive wiki about bacteria
(http://microbewiki.kenyon.edu/index.php/Microbial_Biorealm) and viruses (http://microbewiki.kenyon.edu/index.php
/Viral_Biorealm)
Bacteria that affect crops and other plants (http://www.ncppb.com/)
Bacterial Nomenclature Up-To-Date from DSMZ (http://www.dsmz.de/bactnom/bactname.htm)
Genera of the domain Bacteria (https://web.archive.org/web/20080917230856/http://www.bacterio.cict.fr
/eubacteria.html)—list of Prokaryotic names with Standing in Nomenclature
The largest bacteria (http://www.sciencenews.org/pages/sn_arc99/4_17_99/fob5.htm)
Tree of Life: Eubacteria (http://tolweb.org/tree?group=Eubacteria&contgroup=Life_on_Earth)
Videos (http://www.rowland.harvard.edu/labs/bacteria/index_movies.html) of bacteria swimming and tumbling, use
of optical tweezers and other videos.
Planet of the Bacteria (http://www.stephenjaygould.org/library/gould_bacteria.html) by Stephen Jay Gould
On-line text book on bacteriology (http://www.textbookofbacteriology.net/)
Animated guide to bacterial cell structure. (http://www.blackwellpublishing.com/trun/artwork/Animations/Overview
/overview.html)
Bacteria Make Major Evolutionary Shift in the Lab (https://www.newscientist.com/channel/life/dn14094-bacteria-
make-major-evolutionary-shift-in-the-lab.html)
Online collaboration for bacterial taxonomy. (http://esciencenews.com/articles/2009/02
/19/online.collaboration.identifies.bacteria)
PATRIC (http://patricbrc.org/), a Bioinformatics Resource Center for bacterial pathogens, funded by NIAID
(https://www.niaid.nih.gov/)
Bacterial Chemotaxis Interactive Simulator (http://wormweb.org/bacteriachemo)—A web-app that uses several
simple algorithms to simulate bacterial chemotaxis.
Cell-Cell Communication in Bacteria (https://web.archive.org/web/20090130052842/http://ascb.org/ibioseminars
/bassler/bassler1.cfm) on-line lecture by Bonnie Bassler, and TED: Discovering bacteria's amazing communication
system (http://www.ted.com/index.php/talks/bonnie_bassler_on_how_bacteria_communicate.html)
Sulfur-cycling fossil bacteria from the 1.8-Ga Duck Creek Formation provide promising evidence of evolution's null
hypothesis (http://www.pnas.org/content/early/2015/01/27/1419241112), Proceedings of the National Academy of
Sciences of the United States of America. Summarised in: Scientists discover bacteria that haven't evolved in
more than 2 billion years (http://www.businessinsider.com/scientists-discover-bacteria-that-havent-evolved-in-more-
than-2-billion-years-2015-2), LiveScience and BusinessInsider
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Bacteria

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Bacteria

There are typically 40 million bacterial cells in a gram of soil and a

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Classification and identification

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The ancestors of modern bacteria were unicellular

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Inorganic Organic compounds (lithoheterotrophs) Thermodesulfobacteria,

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Significance in technology and industry

Christian Gottfried Ehrenberg introduced the word "bacterium" in

Antonie van Leeuwenhoek, the first

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